FLÁVIA BADUY VAZ DA SILVA TOLERÂNCIA AMBIENTAL E COMPORTAMENTO SOCIAL E REPRODUTIVO DE Australoheros facetus (CICHLIDAE): QUE MECANISMOS FISIOLÓGICOS E MOLECULARES PERMITEM A ADAPTAÇÃO DESTA ESPÉCIE INVASORA EM PORTUGAL? UNIVERSIDADE DO ALGARVE Faculdade de Ciências e Tecnologia Faro / Portugal 2017 ii FLÁVIA BADUY VAZ DA SILVA TOLERÂNCIA AMBIENTAL E COMPORTAMENTO SOCIAL E REPRODUTIVO DE Australoheros facetus (CICHLIDAE): QUE MECANISMOS FISIOLÓGICOS E MOLECULARES PERMITEM A ADAPTAÇÃO DESTA ESPÉCIE INVASORA EM PORTUGAL? Doutoramento em Ciências Biológicas (Especialidade em Ecofisiologia) Trabalho efetuado sob a orientação de: Prof. Dr. Adelino Vicente Canário Dr. Pedro Miguel G. Guerreiro Dr. João Luis Saraiva UNIVERSIDADE DO ALGARVE Faculdade de Ciências e Tecnologia Faro / Portugal 2017 iii iv Tolerância ambiental e comportamento social e reprodutivo de Australoheros facetus (Cichlidae): que mecanismos fisiológicos e moleculares permitem a adaptação desta espécie invasora em Portugal? Declaração de autoria Declaro ser a autora deste trabalho, que é original e inédito. Autores e trabalhos consultados estão devidamente citados no texto e constam da listagem de referências. I declare to be the author of this work, which is original and unpublished. Authors and works consulted are duly cited in the text and are included in the list of references. ©Flávia Baduy Vaz da Silva, 2017 A Universidade do Algarve tem o direito perpétuo e sem limites geográficos, de arquivar e publicitar este trabalho através de exemplares impressos reproduzidos em papel ou de forma digital, ou por qualquer outro meio conhecido ou que venha a ser inventado, de o divulgar através de repositórios científicos e de admitir a sua cópia e distribuição com objetivos educacionais ou de investigação, não comerciais, desde que seja dado crédito ao autor e editor. This thesis was supported by Brazil's National Council for Scientific and Technological Development (CNPq) through a PhD fellowship from the Science Without Borders Program (245971/2012-2) and by the FCT - Foundation for Science and Technology provided national funds through project UID/Multi/04326/2013. v vi “Todos têm direito a um ambiente de vida humano, sadio e ecologicamente equilibrado e o dever de o defender.” (Artigo 66, Parágrafo 1, da Constituição da República Portuguesa) “Todos têm direito ao meio ambiente ecologicamente equilibrado, bem de uso comum do povo e essencial à sadia qualidade de vida, impondo-se ao poder público e à coletividade o dever de defendê-lo para as presentes e futuras gerações.” (Artigo 225 da Constituição da República Federativa do Brasil) Com meus pés fora da minha zona de conforto! vii viii Acknowledgments First, I’d like to thank to Pedro M. Guerreiro and João L. Saraiva for answering my first e-mail, from far away in Brazil, without recommendations, and thus giving me the chance to develop this project, that changed my life in some many respects. Thank you so much, Pedro and João!!! I really hope that I filled your expectations. Professor Adelino Canário, thank you for allowing me to come to CEIB group at CCMar and become part of the team. It was a great experience, professional and personal. Thank you to all CCMar members (researchers, teachers, students and staff) for receiving me so well. Dear Peter Hubbard… It was so many times that I just turned around and said: “Peeeeeter, sorry, could you help me here….?”. Thanks for be so patient and kind. Let’s be fair: chanchito is much better than tilapia!! I can’t forget to give a really special thanks to Elsa Couto for all support in the lab, especially with the RIA. And, of course, helping me to find everything in the lab! I would be totally lost without you there!! Alex Becker! We got here together… and you have helped me so much in the lab and with sampling… doing some hard work with the filters and tanks… And, talking to me, listening to me when I was angry or upset. Thanks for all the support! Physical and mental! I’d like to thank as well to Sandrinha and Xana. You three together were like my angels in Earth, saving me of feared laboratory. Sandrinha, you know how hard was for me to do the molecular stuff! However, you were there for me, guiding me through the biomolecular world. Xaninha, thank you for all the helping during the field trips and laboratory sampling. I wish you good luck and success in your PhD! I would also like to thank Filipe Ribeiro (FCUL – MARE), the true expert in non-native species, who provided us with valuable information on the distribution of chanchitos, and made available some of the temperature data used in this thesis. Thanks also to Christos Gkenas for funny moments during field trips and conferences! I must not forget to give a special thanks to Carlos Carrapato and colleagues from the Parque Natural do Vale do Guadiana (ICNF-PNVG) for the help during the field work and most importantly, for lending us the electric fishing apparatus, which made collecting the fish possible in the first place. ix Thanks for all the students involved in this thesis, during field trips, fish maintenance, and experiments: Marta Vargas, Jéssica Soares, Mariana Silva, Ana Mariana Fernandes, Joffrey Baeyaert, Louise Merquiol, Diogo Soares, Daniela Teixeira. I’d like to thank for my friends, here and in Brazil, that help me to pass through the PhD without going crazy! Especially to Lica (we can do this!!!), Thiago, Karyna, Nathalia, Nathalie, Cármen and all our coffees! And to my BioGatas from Brazil: Có, Tata, Le, Fer, Iza, Ná, Dani, Penta, Jordes! You are the best!!! Always in my heart! I must give a very special thanks to my cousin-sister, Pamela! I love you so much!!! And I miss you so much!!! Keep betting on MegaSena and come to visit me!!! We have already a lot of things to celebrate together!! Your emotional support was essential to me, to keep me straight in my purposes! Of course, I have to thank for my Family! Mom and Dad and my brother Rafa! I know it was not easy for you to let me go. But you always give me all the support and care that I needed. Always!!! Even in my crazy’s trips or works, months apart from you! And I know that if I have to back home someday, you will be always with open heart and arms! I love you all so much!!!! You are always an example for me! You give me exactly everything that I could need to be a good person! I’m always proud of you! I miss you so much… I wish you could be here to witness my success! If I succeed, was just because of you! // Claro, eu tenho que agradecer pela minha Família! Mãe, Pai e meu irmão Rafa! Eu sei que não foi fácil para vocês me deixarem ir. Mas vocês sempre me deram todo o apoio e cuidado que eu preciso. Sempre!!! Mesmo em minhas viagens ou trabalhos loucos, os quais me separaram durante meses de vocês! E eu sei que, se algum dia eu precisar voltar, vocês sempre estarão com o coração e braços abertos! Eu amo muito todos vocês!!!! Sempre serão um exemplo para mim! Sempre me deram tudo que eu poderia precisar para ser uma boa pessoa! Sinto um imenso orgulho de vocês! Sinto tantas saudades... adoraria que estivessem aqui para testemunhar o meu sucesso! Se eu consegui, foi por causa de vocês! Rui, babe, I have no words to say how important you were during this process! It’s done!!! Thank you for listening to all my complaints, for staying with me when I got sick and taking care of me. Thanks for stay by my side! Thanks for helping me to relax and enjoy Life! This work could not be done without the financial support of CNPq, Science without Borders Program! x Table of contents Acknowledgments ........................................................................................................... ix List of Tables .................................................................................................................. xv List of Figures ............................................................................................................... xvii Resumo ........................................................................................................................ xxiii Abstract ......................................................................................................................... xxv CHAPTER ONE ......................................................................................................... 1 GENERAL INTRODUCTION ..................................................................................... 1 1. Non-native species and Invasion Ecology ............................................................. 3 2. Environmental Tolerance ...................................................................................... 7 2.1. Thermal tolerance ............................................................................................. 10 2.2. Salinity tolerance .............................................................................................. 13 3. Social Behaviour ................................................................................................. 15 3.1. Chemical communication ................................................................................. 16 4. A physiological approach to invasive species ..................................................... 17 5. Target species ...................................................................................................... 19 6. Aim and outline of the thesis ............................................................................... 22 7. References ..........................................................................................................